Leafhoppers (Homoptera: Cicadellidae) that probe human skin...
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© Koninklijke Brill NV, Leiden, 2013 DOI 10.1163/18749836-06001064 Terrestrial Arthropod Reviews 6 (2013) 201–225 brill.com/tar TAR Leafhoppers (Homoptera: Cicadellidae) that probe human skin: A review of the world literature and nineteen new records, from Panama Edwin Domínguez Núñez and Annette Aiello Smithsonian Tropical Research Institute, Apartado 0843-03092 Balboa, Ancon, Republic of Panama e-mails: [email protected]; [email protected] Received on January 16, 2013, Accepted on 17 March 2013. Final version received on April 17, 2013 Summary We review the world literature on leafhoppers (Cicadellidae) that have probed human skin, update the names presented in those published reports, and sort out which are original accounts and which are citations of previous reports (Table 1). We add 19 new records, representing five subfamilies, and at least 13 genera, all from Panama (Table 2). We speculate as to why these normally phytophagous insects sometimes probe human skin, and provide insight into the possible evolutionary consequences of this behavior. Keywords Insecta; Homoptera; Cicadellidae; leafhoppers; aberrant feeding; behavior; bite; blood; entomology; evolution; human skin; probe; medical entomology; new records; Panama Introduction Members of the insect orders Homoptera and Heteroptera, sometimes treated collec- tively as the Hemiptera, have distinctive mouth parts specialized for piercing and suck- ing that have enabled them to radiate into a variety of niches unavailable to members of most other insect groups. Concomitant with this specialization, essentially all Homoptera and Heteroptera subsist on liquid food alone. ough the diets of Heteroptera vary from plant juices, to insect prey, to vertebrate blood, the Homoptera are restricted to plant fluids, mainly the xylem and/or phloem of vascular plants. With their high reproductivity and exceptional capacity to transmit plant diseases, the Homoptera include some of the worst agricultural pests. Among the most important in this respect are the leafhoppers (family Cicadellidae), which include highly efficient vectors of plant viruses (Nault and Rodriguez, 1985; Krinsky, 2002).
Transcript of Leafhoppers (Homoptera: Cicadellidae) that probe human skin...
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© Koninklijke Brill NV, Leiden, 2013 DOI 10.1163/18749836-06001064
Terrestrial Arthropod Reviews 6 (2013) 201–225 brill.com/tarT A R
Leafhoppers (Homoptera: Cicadellidae) that probe human skin: A review of the world literature and nineteen
new records, from Panama
Edwin Domínguez Núñez and Annette Aiello
Smithsonian Tropical Research Institute, Apartado 0843-03092 Balboa, Ancon, Republic of Panama
e-mails: [email protected]; [email protected] Received on January 16, 2013, Accepted on 17 March 2013.
Final version received on April 17, 2013
SummaryWe review the world literature on leafhoppers (Cicadellidae) that have probed human skin, update the names presented in those published reports, and sort out which are original accounts and which are citations of previous reports (Table 1). We add 19 new records, representing five subfamilies, and at least 13 genera, all from Panama (Table 2). We speculate as to why these normally phytophagous insects sometimes probe human skin, and provide insight into the possible evolutionary consequences of this behavior.
KeywordsInsecta; Homoptera; Cicadellidae; leafhoppers; aberrant feeding; behavior; bite; blood; entomology; evolution; human skin; probe; medical entomology; new records; Panama
Introduction
Members of the insect orders Homoptera and Heteroptera, sometimes treated collec-tively as the Hemiptera, have distinctive mouth parts specialized for piercing and suck-ing that have enabled them to radiate into a variety of niches unavailable to members of most other insect groups. Concomitant with this specialization, essentially all Homoptera and Heteroptera subsist on liquid food alone.
Though the diets of Heteroptera vary from plant juices, to insect prey, to vertebrate blood, the Homoptera are restricted to plant fluids, mainly the xylem and/or phloem of vascular plants. With their high reproductivity and exceptional capacity to transmit plant diseases, the Homoptera include some of the worst agricultural pests. Among the most important in this respect are the leafhoppers (family Cicadellidae), which include highly efficient vectors of plant viruses (Nault and Rodriguez, 1985; Krinsky, 2002).
-
202 E.Domínguez Núñez and A. Aiello / Terrestrial Arthropod Reviews 6 (2013) 201–225
Despite their present limitation to feeding on plants, anecdotal evidence suggests that a number of Homoptera, as well as certain plant-feeding Heteroptera, have the potential to diversify onto other food sources. That the switch from herbivory to preda-tion already has occurred within the Heteroptera, e.g., Reduviidae, lends credence to that idea, as does the fact that species representing at least 15 families of herbivorous Heteroptera and five families of Homoptera are known to have bitten humans. These accounts were reviewed by Bequaert (1926), Myers (1929), Usinger (1934), Ryckman (1979), Ryckman and Bentley (1979), and Alexander (1984).
Homoptera known to annoy humans by probing with their mouthparts are found among the leafhoppers (Cicadellidae), treehoppers (Membracidae), spittle bugs (Cercop-idae), plant hoppers (Fulgoroidea), and cicadas (Cicadidae).
Over the years, a variety of non-blood-feeding insects, e.g., lacewing larvae (Neuroptera: Chrysopidae), has “bitten” each of us (EDN and AA) and, independently, we began to collect the leafhoppers involved. A chance conversation led to the joint preparation of this paper, with the objectives of investigating the history of this phenomenon in cicadellids and reporting our own experiences.
Methods
We tracked down every published report of leafhoppers biting humans that we could find, updated the names, and organized the original reports chronologically, each fol-lowed by the publications that cite them. Thus, the entire history of these reports is laid out in Table 1.
Our own observations (Table 2) were made in three general areas of the Republic of Panama: Panama City (immediately east of the Panama Canal); the town of Arraiján (just west of the Canal watershed); and Cocle Province, which is farther west. Attackers were collected manually and preserved in 70% ethanol or were frozen; later, all speci-mens were mounted on points and the male genitalia were preserved in vials in glyc-erin. Vouchers are deposited in MIUP (Museo de Invertebrados G. B. Fairchild de la Universidad de Panamá) and at STRI (Smithsonian Tropical Research Institute) as noted in Table 2.
Nomenclature of the Cicadellidae was updated Cwikla, 1985a; Dietrich and Dmitriev, 2006; Dietrich and Dmitriev, 2008; Dlabola, 1958c; Emeljanov, 1999c; Ghauri, 1971; Ghauri, 1974a; Kramer,1971; Metcalf, 1967a; Metcalf,1967c; Metcalf, 1968a; Ross, 1968; Takiya et al. 2006; Young, 1977.
Results
In all, we found 174 records, published from 1911-2006, by 27 authors (authors and co-authors being counted as one), in 33 publications. These reports represent three cicadellid subfamilies, 19 genera, and at least 27 species from various regions of the world, including China, Cuba, England, French Polynesia, India, Japan, North Africa, the Philippines, Trinidad, and the United States (Table 1). Of the 174 records,
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E.Domínguez Núñez and A. Aiello / Terrestrial Arthropod Reviews 6 (2013) 201–225 203Ta
ble
1. C
icad
ellid
ae re
port
ed to
hav
e pr
obed
hum
an sk
in w
orld
wid
e. O
rigin
al re
port
s are
hig
hlig
hted
and
arr
ange
d ch
rono
logi
cally
, eac
h fo
llow
ed b
y th
e pu
blic
atio
ns th
at c
ite th
em. I
nfor
mat
ion
in sq
uare
bra
cket
s was
add
ed b
y th
e au
thor
s.
Empo
asca
faba
e (H
arris
, 184
1)Ty
phlo
cybi
nae
Dal
las,
Texa
s [U
SA]
Tuck
er (1
911,
p.2
9) (a
s Em
poas
ca m
ali),
spec
imen
s det
erm
ined
by
O
. Hei
dem
ann.
Inge
sted
bloo
d.
Empo
asca
faba
e (H
arris
, 184
1)Ty
phlo
cybi
nae
Rile
y &
Joha
nnse
n (1
915,
p.3
3) (a
s Em
poas
ca m
ali),
sour
ce n
ot
give
n bu
t mos
t lik
ely
Tuck
er (1
911)
.Em
poas
ca fa
bae (
Har
ris, 1
841)
Typh
locy
bina
eLa
wso
n (1
926,
p.7
3), c
iting
Rile
y &
Joha
nnso
n (1
915)
.Em
poas
ca fa
bae (
Har
ris, 1
841)
Typh
locy
bina
eN
orth
Am
eric
aM
yers
(192
9, p
.475
), ci
ting
Tuck
er (1
911)
, Rile
y an
d Jo
hann
sen
(191
5).
Empo
asca
faba
e (H
arris
, 184
1)Ty
phlo
cybi
nae
Ryck
man
& B
entle
y (1
979,
p.3
7-38
) (as
Em
poas
ca m
ali),
citi
ng
Rile
y &
Joha
nnse
n (1
915)
.Em
poas
ca fa
bae (
Har
ris, 1
841)
Typh
locy
bina
eRy
ckm
an &
Ben
tley
(197
9, p
.42)
(as E
mpo
asca
mal
i), c
iting
Tu
cker
(191
1).
Scap
hyto
pius
acu
tus (
Say,
1830
)D
elto
ceph
alin
aeR
iley
& Jo
hann
sen
(191
5, p
.33)
(as P
laty
met
opiu
s acu
tus),
sour
ce
not g
iven
.
Scap
hyto
pius
acu
tus (
Say,
1830
)D
elto
ceph
alin
aeLa
wso
n (1
926,
p.7
3) (a
s Pla
tym
etop
ius a
cutu
s), c
iting
Rile
y &
Jo
hann
sen
(pre
sum
ably
191
5).
Scap
hyto
pius
acu
tus (
Say,
1830
)D
elto
ceph
alin
aeN
orth
Am
eric
aM
yers
(192
9, p
.475
) (as
Pla
tym
etop
ius a
cutu
s), c
iting
Rile
y &
Jo
hann
sen
(191
5).
Scap
hyto
pius
acu
tus (
Say,
1830
)D
elto
ceph
alin
aeR
iley
& Jo
hann
sen
(193
8, p
.143
) (as
Pla
tym
etop
ius a
cutu
s), so
urce
no
t giv
en, b
ut n
ot d
oubt
thei
r ow
n 19
15 re
cord
.Sc
aphy
topi
us a
cutu
s (Sa
y, 18
30)
Del
toce
phal
inae
Ryck
man
& B
entle
y (1
979,
p.3
3) (a
s Pla
tym
etop
ius a
cutu
s), c
iting
M
yers
(192
9).
Scap
hyto
pius
acu
tus (
Say,
1830
)D
elto
ceph
alin
aeRy
ckm
an &
Ben
tley
(197
9, p
.37-
38) (
as P
laty
met
opiu
s acu
tus),
ci
ting
Rile
y &
Joha
nnse
n (1
915)
.
Scap
hyto
pius
acu
tus (
Say,
1830
)D
elto
ceph
alin
aeRy
ckm
an &
Ben
tley
(197
9, p
.38)
(as P
laty
met
opiu
s acu
tus),
citi
ng
Rile
y &
Joha
nnse
n (1
938)
.
(Con
tinue
d )
-
204 E.Domínguez Núñez and A. Aiello / Terrestrial Arthropod Reviews 6 (2013) 201–225
Empo
asca
faba
e (H
arris
, 184
1)Ty
phlo
cybi
nae
Faye
ttevi
lle,
Arka
nsas
[USA
]Be
cker
(191
8, p
.101
) (as
Em
poas
ca m
ali),
spec
imen
s det
erm
ined
by
W.D
. Gib
son.
Empo
asca
faba
e (H
arris
, 184
1)Ty
phlo
cybi
nae
Law
son
(192
0, p
.20)
(as E
mpo
asca
mal
i), c
iting
Bec
ker (
1918
).Em
poas
ca fa
bae (
Har
ris, 1
841)
Typh
locy
bina
eN
orth
Am
eric
aM
yers
(192
9, p
.475
), ci
ting
Beck
er (1
918)
.Em
poas
ca fa
bae (
Har
ris, 1
841)
Typh
locy
bina
eRy
ckm
an (1
979,
p.1
1) (a
s Em
poas
ca m
ali),
citi
ng B
ecke
r (19
18).
Empo
asca
faba
e (H
arris
, 184
1)Ty
phlo
cybi
nae
Ryck
man
& B
entle
y (1
979,
p.3
3), c
iting
Mye
rs (1
929)
.Em
poas
ca fa
bae (
Har
ris, 1
841)
Typh
locy
bina
eAl
exan
der (
1984
, p.5
8) (a
s Em
poas
ca m
ali),
citi
ng B
ecke
r (19
18).
Nep
hote
ttix
nigr
opict
us (S
tål,
1870
)D
elto
ceph
alin
aeM
anila
[Phi
lippi
nes]
Bank
s (19
19, p
.181
) (as
Nep
hote
ttix
apica
lis).
Nep
hote
ttix
nigr
opict
us (S
tål,
1870
)D
elto
ceph
alin
aePh
ilipp
ines
Ryck
man
(197
9, p
.11)
(as N
epho
tetti
x ap
icalis
), ci
ting
Bank
s (1
919)
.N
epho
tetti
x ni
grop
ictus
(Stå
l, 18
70)
Del
toce
phal
inae
Phili
ppin
esAl
exan
der (
1984
, p.5
8) (a
s Nep
hote
ttix
apica
lis),
citin
g Ba
nks
(191
9).
Nep
hote
ttix
vire
scens
(Dist
ant,
1908
)D
elto
ceph
alin
aeM
anila
[Phi
lippi
nes]
Bank
s (19
19, p
.181
) (as
Nep
hote
ttix
bipu
ncta
tus).
Nep
hote
ttix
vire
scens
(Dist
ant,
1908
)D
elto
ceph
alin
aePh
ilipp
ines
Brum
pt (1
922,
p.8
15) (
as N
epho
tetti
x bi
punc
tatu
s), c
iting
Ban
ks
(pre
sum
ably
191
9).
Nep
hote
ttix
vire
scens
(Dist
ant,
1908
)D
elto
ceph
alin
aePh
ilipp
ines
Bequ
aert
(192
6, p
.189
) (as
Nep
hote
ttix
bipu
ncta
tus),
citi
ng B
anks
as
repo
rted
by
Brum
pt (1
922)
.N
epho
tetti
x vi
resce
ns (D
istan
t, 19
08)
Del
toce
phal
inae
Phili
ppin
esBr
umpt
(192
7, p
.952
-953
) (as
Nep
hote
ttix
bipu
ncta
tus),
citi
ng
Bank
s, as
in th
e 19
22 e
ditio
n.N
epho
tetti
x vi
resce
ns (D
istan
t, 19
08)
Del
toce
phal
inae
Phili
ppin
esM
yers
(192
9, p
.474
) (as
Nep
hote
ttix
bipu
ncta
tus),
citi
ng B
rum
pt
(192
2).
Nep
hote
ttix
vire
scens
(Dist
ant,
1908
)D
elto
ceph
alin
aeM
artin
i (19
29, p
.385
) (as
Nep
hote
ttix)
, sou
rce
not g
ive,
but
pr
esum
ably
orig
inat
ing
from
Ban
ks (1
919)
or o
ne o
f tho
se c
iting
hi
m.
Nep
hote
ttix
vire
scens
(Dist
ant,
1908
)D
elto
ceph
alin
aePh
ilipp
ines
Brum
pt (1
936,
p.1
305)
(as N
epho
tetti
x bi
punc
tatu
s), c
iting
Ban
ks,
as in
his
prev
ious
two
editi
ons.
Nep
hote
ttix
vire
scens
(Dist
ant,
1908
)D
elto
ceph
alin
aePh
ilipp
ines
Brum
pt (1
949,
p.1
276)
(as N
epho
tetti
x bi
punc
tatu
s), c
iting
Ban
ks,
as in
his
prev
ious
thre
e ed
ition
s.
Tabl
e 1.
(C
ont.)
(Con
tinue
d )
-
E.Domínguez Núñez and A. Aiello / Terrestrial Arthropod Reviews 6 (2013) 201–225 205
Nep
hote
ttix
vire
scens
(Dist
ant,
1908
)D
elto
ceph
alin
aePh
ilipp
ines
Ryck
man
(197
9, p
.11)
(as N
epho
tetti
x bi
punc
tatu
s), c
iting
Ban
ks
(191
9).
Nep
hote
ttix
vire
scens
(Dist
ant,
1908
)D
elto
ceph
alin
aeRy
ckm
an &
Ben
tley
(197
9, p
.31)
(as N
epho
tetti
x), c
iting
Mar
tini
(192
9).
Nep
hote
ttix
vire
scens
(Dist
ant,
1908
)D
elto
ceph
alin
aeRy
ckm
an &
Ben
tley
(197
9, p
.33)
(as N
epho
tetti
x bi
punc
tatu
s),
citin
g M
yers
(192
9).
Nep
hote
ttix
vire
scens
(Dist
ant,
1908
)D
elto
ceph
alin
aePh
ilipp
ines
Alex
ande
r (19
84, p
.58)
(as N
epho
tetti
x bi
punc
tatu
s, m
isspe
lled
as
bipu
ncta
ta),
citin
g Ba
nks (
1919
).
Empo
asca
faba
e (H
arris
, 184
1)Ty
phlo
cybi
nae
Kan
sas [
USA
]La
wso
n (1
920,
p.2
0) (a
s Em
poas
ca m
ali).
Empo
asca
faba
e (H
arris
, 184
1)Ty
phlo
cybi
nae
Law
son
(192
6, p
.73)
(as E
mpo
asca
mal
i), a
repe
at o
f his
own
repo
rt
of 1
920.
Empo
asca
faba
e (H
arris
, 184
1)Ty
phlo
cybi
nae
Nor
th A
mer
ica
Mye
rs (1
929,
p.4
75),
citin
g La
wso
n (1
926)
.Em
poas
ca fa
bae (
Har
ris, 1
841)
Typh
locy
bina
eR
iley
& Jo
hann
sen
(193
8, p
.143
) (as
Em
poas
ca m
ali),
sour
ce n
ot
give
n.Em
poas
ca fa
bae (
Har
ris, 1
841)
Typh
locy
bina
eK
ansa
s [U
SA]
Ryck
man
& B
entle
y (1
979,
p.2
8), c
iting
Law
son
(192
6).
Empo
asca
faba
e (H
arris
, 184
1)Ty
phlo
cybi
nae
Ryck
man
& B
entle
y (1
979,
p.3
8) (a
s Em
poas
ca m
ali),
citi
ng R
iley
& Jo
hann
sen
(193
8).
Exiti
anus
indi
cus (
Dist
ant,
1908
)D
elto
ceph
alin
aeM
adra
s [In
dia]
Don
ovan
(192
0a, p
.220
; 192
0b, p
.212
) (as
Phr
ynom
orph
us
indi
cus),
bas
ed o
n sp
ecim
ens c
olle
cted
by
him
, tak
en to
Gah
an,
and
dete
rmin
ed b
y D
istan
t.
Exiti
anus
indi
cus (
Dist
ant,
1908
)D
elto
ceph
alin
ael’I
nde
Brum
pt (1
922,
p.8
14) (
as P
hryn
omor
phus
indi
cus),
citi
ng D
onov
an
(pre
sum
ably
192
0).
Exiti
anus
indi
cus (
Dist
ant,
1908
)D
elto
ceph
alin
aeM
adra
s (In
dia)
Berg
evin
(192
5a, p
. 40)
(as A
thys
anus
indi
cus),
citi
ng p
erso
nal
corr
espo
nden
ce fr
om M
arsh
all,
who
repo
rted
that
M.E
. Chi
na
foun
d tw
o sp
ecim
ens i
n th
e Br
itish
Mus
eum
that
wer
e co
llect
ed b
y D
onov
an in
Nov
. 191
9 an
d an
nota
ted
as “
bloo
dsuc
kers
.” U
nkno
wn
to h
im, D
onov
an (1
920)
alre
ady
had
publ
ished
his
acco
unt o
f the
ir bl
ood-
suck
ing
beha
vior
.
(Con
tinue
d )
Tabl
e 1.
(C
ont.)
-
206 E.Domínguez Núñez and A. Aiello / Terrestrial Arthropod Reviews 6 (2013) 201–225
Exiti
anus
indi
cus (
Dist
ant,
1908
)D
elto
ceph
alin
aeM
adra
s, In
dia
Bequ
aert
(192
6, p
.189
) (as
Phr
ynom
orph
us in
dicu
s), c
iting
D
onov
an (1
920)
.Ex
itian
us in
dicu
s (D
istan
t, 19
08)
Del
toce
phal
inae
Indi
aBr
umpt
(192
7, p
.952
) (as
Phr
ynom
orph
us in
dicu
s), c
iting
Don
ovan
(p
resu
mab
ly 1
920)
.Ex
itian
us in
dicu
s (D
istan
t, 19
08)
Del
toce
phal
inae
Indi
aM
yers
(192
9, p
.474
) [as
Ath
ysan
us (P
hryn
omor
phus
) ind
icus],
citi
ng
Don
ovan
(192
0).
Exiti
anus
indi
cus (
Dist
ant,
1908
)D
elto
ceph
alin
aeM
adra
s (In
dia)
Chi
na (1
929,
p.9
7) (a
s Ath
ysan
us in
dicu
s), in
not
e re
ad b
y E.
B.
Poul
ton
at 1
6 O
ct. 1
929
mee
ting
of th
e En
tom
olog
ical
Soc
iety
of
Lond
on, i
n w
hich
he
men
tions
that
he
“dre
w B
erge
vin’s
atte
ntio
n”
to D
onov
an sp
ecim
ens i
n th
e Br
itish
Mus
eum
.Ex
itian
us in
dicu
s (D
istan
t, 19
08)
Del
toce
phal
inae
Indi
aBr
umpt
(193
6, p
.130
5) (a
s Phr
ynom
orph
us in
dicu
s), c
iting
D
onov
an (p
resu
mab
ly 1
920)
, as i
n hi
s 192
7 ed
ition
.Ex
itian
us in
dicu
s (D
istan
t, 19
08)
Del
toce
phal
inae
Mad
ras,
Inde
an
glai
seFo
ley
& A
udou
ze (1
938,
p.5
26) (
as A
thys
anus
indi
cus),
repo
rtin
g ob
serv
atio
ns b
y D
onov
an in
Nov
. 191
9.Ex
itian
us in
dicu
s (D
istan
t, 19
08)
Del
toce
phal
inae
Indi
aBr
umpt
(194
9, p
.127
5) (a
s Ath
ysan
us in
dicu
s), c
iting
Don
ovan
(p
resu
mab
ly 1
920)
, as i
n hi
s pre
viou
s tw
o ed
ition
s.Ex
itian
us in
dicu
s (D
istan
t, 19
08)
Del
toce
phal
inae
Mad
ras,
Indi
aRy
ckm
an (1
979,
p.1
6) (a
s Phr
ynom
orph
us in
dicu
s), c
iting
Don
ovan
(1
920a
).Ex
itian
us in
dicu
s (D
istan
t, 19
08)
Del
toce
phal
inae
Ryck
man
& B
entle
y (1
979,
p.3
3) (a
s Ath
ysan
us in
dicu
s), c
iting
M
yers
(192
9).
Nep
hote
ttix
vire
scens
(Dist
ant,
1908
)D
elto
ceph
alin
aeC
alcu
tta [I
ndia
]Br
umpt
(192
2, p
.815
) (as
Nep
hote
ttix
bipu
ncta
tus),
citi
ng A
. Al
cock
, with
no
date
or p
lace
of p
ublic
atio
n. W
e ha
ve b
een
unab
le
to lo
cate
any
Alc
ock
publ
icat
ion
that
men
tions
Cic
adel
lidae
as
bitin
g hu
man
s.
Nep
hote
ttix
vire
scens
(Dist
ant,
1908
)D
elto
ceph
alin
aeC
alcu
tta [I
ndia
]Be
quae
rt (1
926,
p.1
89) (
as N
epho
tetti
x bi
punc
tatu
s), c
iting
Alc
ock
as
repo
rted
in B
rum
pt (1
922)
.N
epho
tetti
x vi
resce
ns (D
istan
t, 19
08)
Del
toce
phal
inae
Cal
cutta
(Ind
ia)
Brum
pt (1
927,
p.9
52-9
53) (
as N
epho
tetti
x bi
punc
tatu
s), c
iting
A.
Alc
ock,
as i
n th
e 19
22 e
ditio
n.N
epho
tetti
x vi
resce
ns (D
istan
t, 19
08)
Del
toce
phal
inae
Indi
aM
yers
(192
9, p
.474
) (as
Nep
hotet
tix b
ipun
ctatu
s), c
iting
Bru
mpt
(1
922)
.
Tabl
e 1.
(C
ont.)
(Con
tinue
d )
-
E.Domínguez Núñez and A. Aiello / Terrestrial Arthropod Reviews 6 (2013) 201–225 207
Nep
hote
ttix
vire
scens
(Dist
ant,
1908
)D
elto
ceph
alin
aeC
alcu
tta (I
ndia
)Br
umpt
(193
6, p
.130
5) (a
s Nep
hote
ttix
bipu
ncta
tus),
citi
ng A
. Al
cock
, as i
n hi
s pre
viou
s 2 e
ditio
ns.
Nep
hote
ttix
vire
scens
(Dist
ant,
1908
)D
elto
ceph
alin
aeC
alcu
tta (I
ndia
)Br
umpt
(194
9, p
.127
6) (a
s Nep
hote
ttix
bipu
ncta
tus),
citi
ng A
. Al
cock
(miss
pelle
d as
Alc
ok).
Exiti
anus
fasci
olat
us (M
elic
har,
1911
)D
elto
ceph
alin
aeIn
Sal
ah [s
outh
ern
Alge
ria]
Berg
evin
(192
5a, p
.38-
39) (
as A
thys
anus
und
escr
ibed
sp.);
de
scrib
ed b
y hi
m in
a fo
llow
ing
pape
r (19
25b,
p.4
2) a
s Ath
ysan
us
vuln
eran
s, fro
m si
x sp
ecim
ens (
4 m
ales
, 2 fe
mal
es) s
ent t
o hi
m b
y D
r. Fo
ley,
who
repo
rted
thei
r biti
ng b
ehav
ior a
s com
mun
icat
ed in
a
lette
r fro
m D
r. Fo
uque
, dat
ed 1
2 Se
pt. 1
924.
Exiti
anus
fasci
olat
us (M
elic
har,
1911
)D
elto
ceph
alin
aeIn
Sal
ah (s
outh
ern
Alge
ria)
Berg
evin
(192
5b, p
p. 4
2) (a
s Ath
ysan
us v
ulne
rans
sp. n
ov.),
from
sp
ecim
ens c
olle
cted
by
Dr.
Fouq
ue, a
nd se
nt to
him
by
Dr.
Fole
y.Ex
itian
us fa
sciol
atus
(Mel
icha
r, 19
11)
Del
toce
phal
inae
“Sah
ara”
in e
rror
Bequ
aert
(192
6, p
.189
) (as
Ath
ysan
us v
ulne
rans
), ci
ting
Berg
evin
(1
925)
, but
he
misr
ead
Sala
h as
“Sa
hara
.”Ex
itian
us fa
sciol
atus
(Mel
icha
r, 19
11)
Del
toce
phal
inae
“Sah
ara”
in e
rror
Mye
rs (1
929,
p.4
74) (
as A
thys
anus
vul
nera
ns),
citin
g Be
rgev
in
(192
5b),
thou
gh p
erpe
tuat
ing
Bequ
aert’
s err
or c
once
rnin
g th
e lo
catio
n; p
erha
ps h
e di
d no
t hav
e ac
cess
to B
erge
vin’s
pub
licat
ion.
Exiti
anus
fasci
olat
us (M
elic
har,
1911
)D
elto
ceph
alin
aeIn
Sal
ah (s
outh
ern
Alge
ria)
Chi
na (1
929,
p.9
7) (a
s Ath
ysan
us v
ulne
rans
), ci
ting
Berg
evin
(1
925a
& 1
925b
), in
lette
r rea
d by
E.B
. Pou
lton
at 1
6 O
ct. 1
929
mee
ting
of th
e En
tom
olog
ical
Soc
iety
of L
ondo
n.Ex
itian
usD
elto
ceph
alin
aeM
artin
i (19
29, p
.385
) (as
Ath
ysanu
s), so
urce
not
giv
en, b
ut b
ased
on
the m
atch
ing
list o
f gen
era,
pre
sum
ably
orig
inat
ing
from
Be
rgev
in (1
925a
).Ex
itian
us fa
sciol
atus
(Mel
icha
r, 19
11)
Del
toce
phal
inae
Alge
riaRy
ckm
an (1
979,
p.1
2) (a
s Ath
ysan
us sp
.), c
iting
Ber
gevi
n (1
925)
.Ex
itian
usD
elto
ceph
alin
aeRy
ckm
an &
Ben
tley
(197
9, p
.31)
(as A
thys
anus
sp.),
citi
ng M
artin
i (1
929)
.Ex
itian
us fa
sciol
atus
(Mel
icha
r, 19
11)
Del
toce
phal
inae
Ryck
man
& B
entle
y (1
979,
p.3
3) (a
s Ath
ysan
us v
ulne
rans
), ci
ting
Mye
rs (1
929)
.
(Con
tinue
d )
Tabl
e 1.
(C
ont.)
-
208 E.Domínguez Núñez and A. Aiello / Terrestrial Arthropod Reviews 6 (2013) 201–225
Eusce
lis cu
rtice
ps L
indb
erg,
192
7D
elto
ceph
alin
aeK
hart
oum
(Sou
dan
Egyp
tien)
Berg
evin
(192
5a, p
p. 3
9-40
) (as
Ath
ysan
us sp
.), re
gard
ing
a sp
ecim
en c
olle
cted
by
H.B
. Joh
nsto
n, 1
7 O
ct. 1
924,
and
sent
by
Mar
shal
l to
Berg
evin
, who
pas
sed
it on
to L
indb
erg
who
then
de
scrib
ed it
(192
7, p
.92-
94) a
s Eus
celis
curt
iceps
.
Eusce
lis cu
rtice
ps L
indb
erg,
192
7D
elto
ceph
alin
aeK
hart
oum
, Ang
lo-
Egyp
tian
Suda
nBe
quae
rt (1
926,
p. 1
89) (
as A
thys
anus
), ci
ting
Berg
evin
(192
5).
Eusce
lis cu
rtice
ps L
indb
erg,
192
7D
elto
ceph
alin
aeK
hart
oum
(Gou
ver-
nem
ent d
u So
udan
)Li
ndbe
rg (1
927c
, p. 9
2), d
escr
ibin
g sp
ecim
ens s
ent t
o hi
m b
y Be
rgev
in.
Eusce
lis cu
rtice
ps L
indb
erg,
192
7D
elto
ceph
alin
aeSu
dan
Mye
rs (1
929,
p.4
75),
citin
g Li
ndbe
rg (1
927)
.Eu
scelis
curt
iceps
Lin
dber
g, 1
927
Del
toce
phal
inae
Suda
nC
hina
(192
9, p
.97)
, citi
ng B
erge
vin
(192
5a),
in le
tter r
ead
by
Poul
ton
at 1
6 O
ct. 1
929
mee
ting
of th
e En
tom
olog
ical
Soc
iety
of
Lond
on.
Eusce
lis cu
rtice
ps L
indb
erg,
192
7D
elto
ceph
alin
aeK
hart
oum
(Sud
an)
Fole
y &
Aud
ouze
(193
8, p
.526
) (as
Ath
ysan
us),
men
tioni
ng th
at
Mar
shal
l sen
t 5 u
ndes
crib
ed sp
ecie
s of b
lood
-suc
king
“Ja
ssid
ae”
to
Berg
evin
in 1
925.
Eusce
lis cu
rtice
ps L
indb
erg,
192
7D
elto
ceph
alin
aeSu
dan
Ryck
man
& B
entle
y (1
979,
p.2
8), c
iting
Lin
dber
g (1
927)
.Eu
scelis
curt
iceps
Lin
dber
g, 1
927
Del
toce
phal
inae
Kha
rtou
m (S
udan
)Le
wis
(195
8, p
.45)
(miss
pelle
d as
curt
ipes)
, citi
ng L
indb
erg
(192
7).
Eusce
lis cu
rtice
ps L
indb
erg,
192
7D
elto
ceph
alin
aeSu
dan
Ryck
man
& B
entle
y (1
979,
p.2
8), c
iting
Lew
is (1
958)
.Eu
scelis
curt
iceps
Lin
dber
g, 1
927
Del
toce
phal
inae
Ryck
man
& B
entle
y (1
979,
p.3
3), c
iting
Mye
rs (1
929)
.Eu
scelis
curt
iceps
Lin
dber
g, 1
927
Del
toce
phal
inae
Nor
th A
frica
Alex
ande
r (19
84, p
.58)
(miss
pelle
d as
Ath
ysau
rus s
p.),
citin
g Be
rgev
in (1
925)
.
Acon
urell
a ne
ar p
rolix
a (L
ethi
erry
, 188
5)D
elto
ceph
alin
aeK
hart
oum
(Sou
dan
Egyp
tien)
Berg
evin
(192
5a, p
p. 3
9-40
) (as
Tham
note
ttix,
miss
pelle
d as
Th
amot
ettiy
), re
gard
ing
spec
imen
s col
lect
ed b
y H
.B. J
ohns
ton,
17
Oct
. 192
4, a
nd se
nt b
y M
arsh
all t
o Be
rgev
in, w
ho p
asse
d th
em o
n to
Lin
dber
g w
ho th
en d
escr
ibed
them
(192
7, p
.88-
90) a
s 5
spec
imen
s of Th
amno
tetti
x sa
ngui
suga
. Lat
er, M
yers
(192
9, p
.474
) re
port
ed th
at a
ccor
ding
to W
. E. C
hina
, the
y co
rrec
tly b
elon
g to
Ac
onur
a.
Tabl
e 1.
(C
ont.)
(Con
tinue
d )
-
E.Domínguez Núñez and A. Aiello / Terrestrial Arthropod Reviews 6 (2013) 201–225 209
Acon
urell
a ne
ar p
rolix
a (L
ethi
erry
, 188
5)D
elto
ceph
alin
aeK
hart
oum
, Ang
lo-
Egyp
tian
Suda
nBe
quae
rt (1
926,
p. 1
89) (
as Th
amno
tetti
x), c
iting
Ber
gevi
n (1
925)
.
Acon
urell
a ne
ar p
rolix
a (L
ethi
erry
, 188
5)D
elto
ceph
alin
aeK
hart
oum
(Gou
ver-
nem
ent d
u So
udan
)Li
ndbe
rg (1
927c
, p. 8
8), d
escr
ibin
g (a
s Tham
note
ttix
sang
uisu
ga)
spec
imen
s sen
t to
him
by
Berg
evin
. Lat
er, W
. E. C
hina
said
they
co
rrec
tly b
elon
g to
Aco
nura
.Ac
onur
ella
near
pro
lixa
(Let
hier
ry, 1
885)
Del
toce
phal
inae
Suda
nM
yers
(192
9, p
.474
) (as
Tham
note
ttix
sang
uisu
ga),
citin
g Li
ndbe
rg
(192
7), w
ho d
escr
ibed
it a
s a n
ew sp
ecie
s, Th
amno
tetti
x sa
ngui
suga
, bu
t in
foot
note
, Mye
rs sa
ys W
.E. C
hina
ass
ured
him
it b
elon
gs to
Ac
onur
a ve
ry n
ear A
. pro
lixa.
Acon
urell
a ne
ar p
rolix
a (L
ethi
erry
, 188
5)D
elto
ceph
alin
aeSu
dan
Chi
na (1
929,
p.9
7) (a
s Tham
note
ttix
sang
uisu
ga),
citin
g Be
rgev
in
(192
5a),
in le
tter r
ead
by E
.B. P
oulto
n at
16
Oct
ober
192
9 m
eetin
g of
the
Ento
mol
gica
l Soc
iety
of L
ondo
n.Ac
onur
ella
near
pro
lixa
(Let
hier
ry, 1
885)
Del
toce
phal
inae
Mar
tini (
1929
, p.3
85) (
as Th
amno
tetti
x), s
ourc
e no
t giv
en, b
ut
base
d on
the
mat
chin
g lis
t of g
ener
a, p
resu
mab
ly o
rigin
atin
g fro
m
Berg
evin
(192
5a).
Acon
urell
a ne
ar p
rolix
a (L
ethi
erry
, 188
5)D
elto
ceph
alin
aeK
hart
oum
(Sud
an)
Fole
y &
Aud
ouze
(193
8, p
.526
) (as
Ath
ysan
us),
men
tioni
ng th
at
Mar
shal
l sen
t 5 u
ndes
crib
ed sp
ecie
s of b
lood
-suc
king
“Ja
ssid
ae”
to
Berg
evin
in 1
925.
Acon
urell
a ne
ar p
rolix
a (L
ethi
erry
, 188
5)D
elto
ceph
alin
aeBr
umpt
(194
9, p
.127
6) (a
s Tham
note
ttix,
miss
pelle
d as
Ta
mno
tetti
x), c
iting
Ber
gevi
n (1
925)
.Ac
onur
ella
near
pro
lixa
(Let
hier
ry, 1
885)
Del
toce
phal
inae
Kha
rtou
m (S
udan
)Le
wis
(195
8, p
.45)
(as A
conu
ra n
r. pr
olix
a: a
s “a
bug
whi
ch m
ay b
e an
Aco
nura
”), c
iting
Lin
dber
g (1
927)
and
Mye
rs (1
929)
.Ac
onur
ella
near
pro
lixa
(Let
hier
ry, 1
885)
Del
toce
phal
inae
Suda
nRy
ckm
an &
Ben
tley
(197
9, p
.28)
(as Th
amno
tetti
x sa
ngui
suga
), ci
ting
Lind
berg
(192
7).
Acon
urell
a ne
ar p
rolix
a (L
ethi
erry
, 188
5)D
elto
ceph
alin
aeRy
ckm
an &
Ben
tley
(197
9, p
.33)
(as Th
amno
tetti
x sa
ngui
suga
), ci
ting
Mye
rs (1
929)
.Ac
onur
ella
near
pro
lixa
(Let
hier
ry, 1
885)
Del
toce
phal
inae
Ryck
man
& B
entle
y (1
979,
p.3
1) (a
s Tham
note
ttix)
, citi
ng M
artin
i (1
929)
.
(Con
tinue
d )
Tabl
e 1.
(C
ont.)
-
210 E.Domínguez Núñez and A. Aiello / Terrestrial Arthropod Reviews 6 (2013) 201–225
Oro
sius c
ellul
osus
(Lin
dber
g, 1
927)
Del
toce
phal
inae
Kha
rtou
m (S
ouda
n Eg
yptie
n)Be
rgev
in (1
925a
, pp.
39-
40) (
as D
eltoc
epha
lus),
rega
rdin
g sp
eci-
men
s col
lect
ed b
y H
.B. J
ohns
ton,
17
Oct
. 192
4, a
nd se
nt b
y M
arsh
all t
o Be
rgev
in, w
ho p
asse
d th
em o
n to
Lin
dber
g, w
ho
desc
ribed
them
(192
7, p
.90-
92) a
s thr
ee sp
ecim
ens o
f Tham
note
ttix
cellu
losu
s (m
isspe
lled
as ce
llulo
sa).
Oro
sius c
ellul
osus
(Lin
dber
g, 1
927)
Del
toce
phal
inae
Kha
rtou
m, A
nglo
- Eg
yptia
n Su
dan
Bequ
aert
(192
6, p
. 189
) (as
Delt
ocep
halu
s), c
iting
Ber
gevi
n (1
925)
.
Oro
sius c
ellul
osus
(Lin
dber
g, 1
927)
Del
toce
phal
inae
Kha
rtou
m (G
ouve
r-ne
men
t du
Soud
an)
Lind
berg
(192
7c, p
. 90)
, des
crib
ing
(as Th
amno
tetti
x ce
llulo
sa) a
sp
ecim
en se
nt to
him
by
Berg
evin
.O
rosiu
s cell
ulos
us (L
indb
erg,
192
7)D
elto
ceph
alin
aeSu
dan
Mye
rs (1
929,
p.4
75) (
as Th
amno
tetti
x ce
llulo
sa),
citin
g Li
ndbe
rg
(192
7).
Oro
sius c
ellul
osus
(Lin
dber
g, 1
927)
Del
toce
phal
inae
Suda
nC
hina
(192
9, p
.97)
(as Th
amno
tetti
x ce
llulo
sa),
citin
g Be
rgev
in
(192
5a),
in le
tter r
ead
by E
.B. P
oulto
n at
16
Oct
. 192
9 m
eetin
g of
th
e En
tom
olog
ical
Soc
iety
of L
ondo
n.O
rosiu
s cell
ulos
us (L
indb
erg,
192
7)D
elto
ceph
alin
aeM
artin
i (19
29, p
.385
) (as
Delt
ocep
halu
s), so
urce
not
giv
en, b
ut
base
d on
the
mat
chin
g lis
t of g
ener
a, p
resu
mab
ly o
rigin
atin
g fro
m
Berg
evin
(192
5a).
Oro
sius c
ellul
osus
(Lin
dber
g, 1
927)
Del
toce
phal
inae
Brum
pt (1
949,
p.1
276)
(as D
eltoc
epha
lus),
citi
ng B
erge
vin
(192
5).
Oro
sius c
ellul
osus
(Lin
dber
g, 1
927)
Del
toce
phal
inae
Kha
rtou
m (S
udan
)Le
wis
(195
8, p
.45-
46) (
as Th
amno
tetti
x ce
llulo
sa),
citin
g Li
ndbe
rg
(192
7).
Oro
sius c
ellul
osus
(Lin
dber
g, 1
927)
Del
toce
phal
inae
Suda
nRy
ckm
an &
Ben
tley
(197
9, p
.28)
(as Th
amno
tetti
x ce
llulo
sa),
citin
g Li
ndbe
rg (1
927)
.O
rosiu
s cell
ulos
us (L
indb
erg,
192
7)D
elto
ceph
alin
aeRy
ckm
an &
Ben
tley
(197
9, p
.28)
(as Th
amno
tetti
x ce
llulo
sa),
citin
g Le
wis
(195
8).
Oro
sius c
ellul
osus
(Lin
dber
g, 1
927)
Del
toce
phal
inae
Ryck
man
& B
entle
y (1
979,
p.3
1) (a
s Delt
ocep
halu
s), c
iting
Mar
tini
(192
9).
Oro
sius c
ellul
osus
(Lin
dber
g, 1
927)
Del
toce
phal
inae
Ryck
man
& B
entle
y (1
979,
p.3
3) (a
s Tham
note
ttix
cellu
losa
), ci
ting
Mye
rs (1
929)
.
Erat
oneu
ra b
asila
ris (S
ay, 1
825)
Typh
locy
bina
eLa
wso
n (1
926,
p.7
3) (a
s Ery
thro
neur
a ba
silar
is), c
iting
unp
ublis
hed
expe
rienc
e of
Ray
mon
d H
. Bea
mer
.
Tabl
e 1.
(C
ont.)
(Con
tinue
d )
-
E.Domínguez Núñez and A. Aiello / Terrestrial Arthropod Reviews 6 (2013) 201–225 211
Erat
oneu
ra b
asila
ris (S
ay, 1
825)
Typh
locy
bina
eM
yers
(192
9, p
.475
) (as
Ery
thro
neur
a ba
silar
is), c
iting
Law
son
(192
6).
Erat
oneu
ra b
asila
ris (S
ay, 1
825)
Typh
locy
bina
eR
iley
& Jo
hann
sen
(193
8, p
.143
) (Er
ythr
oneu
ra b
asila
ris),
citin
g La
wso
n (1
926)
.Er
aton
eura
bas
ilaris
(Say
, 182
5)Ty
phlo
cybi
nae
Kan
sas [
USA
]Ry
ckm
an &
Ben
tley
(197
9, p
.28)
(as E
ryth
rone
ura
basil
aris)
, citi
ng
Law
son
(192
6).
Erat
oneu
ra b
asila
ris (S
ay, 1
825)
Typh
locy
bina
eRy
ckm
an &
Ben
tley
(197
9, p
.33)
(as E
ryth
rone
ura
basil
aris)
, citi
ng
Mye
rs (1
929)
.Er
aton
eura
bas
ilaris
(Say
, 182
5)Ty
phlo
cybi
nae
Ryck
man
& B
entle
y (1
979,
p.3
8) (a
s Ery
thro
neur
a ba
silar
is), c
iting
R
iley
& Jo
hann
sen
(193
8).
Para
phlep
sius i
rror
atus
(Say
, 183
0)D
elto
ceph
alin
aeLa
wso
n (1
926,
p.7
4) (a
s Phl
epsiu
s irr
orat
us),
citin
g un
publ
ished
ex
perie
nce
of E
.M. B
ecto
n.
Para
phlep
sius i
rror
atus
(Say
, 183
0)D
elto
ceph
alin
aeN
orth
Am
eric
aM
yers
(192
9, p
.475
) (as
Phl
epsiu
s irr
orat
us),
citin
g La
wso
n (1
926)
.Pa
raph
lepsiu
s irr
orat
us (S
ay, 1
830)
Del
toce
phal
inae
Rile
y &
Joha
nnse
n (1
938,
p.1
43) (
as P
hlep
sius i
rror
atus
), ci
ting
Law
son
(192
6).
Para
phlep
sius i
rror
atus
(Say
, 183
0)D
elto
ceph
alin
aeK
ansa
s [U
SA]
Ryck
man
& B
entle
y (1
979,
p.2
8) (a
s Phl
epsiu
s irr
orat
us),
citin
g La
wso
n (1
926)
.Pa
raph
lepsiu
s irr
orat
us (S
ay, 1
830)
Del
toce
phal
inae
Ryck
man
& B
entle
y (1
979,
p.3
3) (a
s Phl
epsiu
s irr
orat
us),
citin
g M
yers
(192
9).
Para
phlep
sius i
rror
atus
(Say
, 183
0)D
elto
ceph
alin
aeRy
ckm
an &
Ben
tley
(197
9, p
.38)
(as P
hlep
sius i
rror
atus
), ci
ting
Rile
y &
Joha
nnse
n (1
938)
.
Endr
ia in
imica
(Say
, 183
0)D
elto
ceph
alin
aeLa
wso
n (1
926,
p.7
4) (a
s Delt
ocep
halu
s ini
micu
s).
Endr
ia in
imica
(Say
, 183
0)D
elto
ceph
alin
aeN
orth
Am
eric
aM
yers
(192
9, p
.475
) (as
Delt
ocep
halu
s ini
micu
s), c
iting
Law
son
(192
6).
Endr
ia in
imica
(Say
, 183
0)D
elto
ceph
alin
aeR
iley
& Jo
hann
sen
(193
8, p
.143
) (as
Delt
ocep
halu
s ini
micu
s),
citin
g La
wso
n (1
926)
.En
dria
inim
ica (S
ay, 1
830)
Del
toce
phal
inae
Kan
sas [
USA
]Ry
ckm
an &
Ben
tley
(197
9, p
.28)
(as D
eltoc
epha
lus i
nim
icus),
ci
ting
Law
son
(192
6).
(Con
tinue
d )
Tabl
e 1.
(C
ont.)
-
212 E.Domínguez Núñez and A. Aiello / Terrestrial Arthropod Reviews 6 (2013) 201–225
Endr
ia in
imica
(Say
, 183
0)D
elto
ceph
alin
aeRy
ckm
an &
Ben
tley
(197
9, p
.33)
(as D
eltoc
epha
lus i
nim
icus),
ci
ting
Mye
rs (1
929)
.En
dria
inim
ica (S
ay, 1
830)
Del
toce
phal
inae
Ryck
man
& B
entle
y (1
979,
p.3
8) (a
s Delt
ocep
halu
s ini
micu
s),
citin
g R
iley
& Jo
hann
sen
(193
8).
Dra
ecul
acep
hala
mol
lipes
(Say
, 184
0)C
icad
ellin
aeLa
wso
n (1
926,
p.7
4).
Dra
ecul
acep
hala
mol
lipes
(Say
, 184
0)C
icad
ellin
aeN
orth
Am
eric
aM
yers
(192
9, p
.474
) (m
isspe
lled
as D
racu
laec
epha
la m
ollip
es),
citin
g La
wso
n (1
926)
.D
raec
ulac
epha
la m
ollip
es (S
ay, 1
840)
Cic
adel
linae
Rile
y &
Joha
nnse
n (1
938,
p.1
43),
citin
g La
wso
n (1
926)
.D
raec
ulac
epha
la m
ollip
es (S
ay, 1
840)
Cic
adel
linae
Kan
sas [
USA
]Ry
ckm
an &
Ben
tley
(197
9, p
.28)
, citi
ng L
awso
n (1
926)
.D
raec
ulac
epha
la m
ollip
es (S
ay, 1
840)
Cic
adel
linae
Ryck
man
& B
entle
y (1
979,
p.3
3), c
iting
Mye
rs (1
929)
.D
raec
ulac
epha
la m
ollip
es (S
ay, 1
840)
Cic
adel
linae
Ryck
man
& B
entle
y (1
979,
p.3
8), c
iting
Rile
y &
Joha
nnse
n (1
938)
.
Exiti
anus
obs
curin
ervi
s (St
ål, 1
859)
Del
toce
phal
inae
Law
son
(192
6, p
.74)
(as E
usce
lis o
bscu
riner
vis).
Exiti
anus
obs
curin
ervi
s (St
ål, 1
859)
Del
toce
phal
inae
Nor
th A
mer
ica
Mye
rs (1
929,
p.4
75) (
as E
usce
lis o
bscu
riner
vis),
citi
ng L
awso
n (1
926)
.Ex
itian
us o
bscu
riner
vis (
Stål
, 185
9)D
elto
ceph
alin
aeR
iley
& Jo
hann
sen
(193
8, p
.143
) (as
Eus
celis
obs
curin
ervi
s), c
iting
La
wso
n (1
926)
.Ex
itian
us o
bscu
riner
vis (
Stål
, 185
9)D
elto
ceph
alin
aeK
ansa
s [U
SA]
Ryck
man
& B
entle
y (1
979,
p.2
8) (E
usce
lis o
bscu
riner
vis),
citi
ng
Law
son
(192
6).
Exiti
anus
obs
curin
ervi
s (St
ål, 1
859)
Del
toce
phal
inae
Ryck
man
& B
entle
y (1
979,
p.3
3) (a
s Eus
celis
obs
curin
ervi
s), c
iting
M
yers
(192
9).
Exiti
anus
obs
curin
ervi
s (St
ål, 1
859)
Del
toce
phal
inae
Ryck
man
& B
entle
y (1
979,
p.3
8) (a
s Eus
celis
obs
curin
ervi
s), c
iting
R
iley
& Jo
hann
sen
(193
8).
Psam
mot
ettix
livi
dellu
s (Ze
tters
tedt
, 18
40) o
r nea
rD
elto
ceph
alin
aeH
such
owfu
, K
iang
su, C
hina
Law
son
(192
6, p
.74)
(as D
eltoc
epha
lus s
triat
us o
r nea
r), c
iting
co
rres
pond
ence
from
Mar
shal
Her
tig.
Psam
mot
ettix
livi
dellu
s (Ze
tters
tedt
, 18
40) o
r nea
rD
elto
ceph
alin
aeK
iang
su, C
hina
Mye
rs (1
929,
p.4
75) (
as D
eltoc
epha
lus),
citi
ng L
awso
n (1
926)
.
Tabl
e 1.
(C
ont.)
(Con
tinue
d )
-
E.Domínguez Núñez and A. Aiello / Terrestrial Arthropod Reviews 6 (2013) 201–225 213
Psam
mot
ettix
livi
dellu
s (Ze
tters
tedt
, 18
40) o
r nea
rD
elto
ceph
alin
aeK
iang
su, C
hina
Ryck
man
& B
entle
y (1
979,
p.2
8) (a
s Delt
ocep
halu
s), c
iting
Law
son
(192
6).
Psam
mot
ettix
livi
dellu
s (Ze
tters
tedt
, 18
40) o
r nea
rD
elto
ceph
alin
aeRy
ckm
an &
Ben
tley
(197
9, p
.33)
(as D
eltoc
epha
lus),
citi
ng M
yers
(1
929)
.
Exiti
anus
exiti
osus
(Uhl
er, 1
880)
Del
toce
phal
inae
St. L
ouis,
Miss
ouri
[USA
]C
rosb
y (1
926,
p.8
12) (
as E
usce
lis ex
itios
us),
dete
rmin
ed b
y W
.L.
McA
tee.
Exiti
anus
exiti
osus
(Uhl
er, 1
880)
Del
toce
phal
inae
Nor
th A
mer
ica
Mye
rs (1
929,
p.4
75) (
as E
usce
lis ex
itios
us),
citin
g C
rosb
y (1
926)
.Ex
itian
us ex
itios
us (U
hler
, 188
0)D
elto
ceph
alin
aeSt
. Lou
is, M
issou
ri [U
SA]
Ryck
man
(197
9, p
.15)
(as E
usce
lis ex
itios
us),
citin
g C
rosb
y (1
926)
.
Exiti
anus
exiti
osus
(Uhl
er, 1
880)
Del
toce
phal
inae
Ryck
man
& B
entle
y (1
979,
p.3
3) (a
s Eus
celis
exiti
osus
), ci
ting
Mye
rs (1
929)
.Ex
itian
us ex
itios
us (U
hler
, 188
0)D
elto
ceph
alin
aeU
SAAl
exan
der (
1984
, p.5
8) (a
s Eus
celis
exiti
osus
), ci
ting
Cro
sby
(192
6).
Empo
asca
faba
e (H
arris
, 184
1), a
false
re
cord
Typh
locy
bina
eRe
adio
(192
7) p
ublis
hed
“Stu
dies
on
the
biol
ogy
of th
e Re
duvi
idae
of
Am
eric
a N
orth
of M
exic
o,”
but a
s far
as w
e ca
n de
term
ine
didn
’t m
entio
n C
icad
ellid
ae.
Empo
asca
faba
e (H
arris
, 184
1), a
false
re
cord
Typh
locy
bina
eRy
ckm
an &
Ben
tley
(197
9, p
.36)
(as E
mpo
asca
mal
i), c
iting
Re
adio
(192
7) in
err
or; R
eadi
o ne
ver p
ublis
hed
on b
iting
ci
cade
llids
.Em
poas
ca fa
bae (
Har
ris, 1
841)
, a fa
lse
reco
rdTy
phlo
cybi
nae
Alex
ande
r (19
84, p
.58)
(as E
mpo
asca
mal
i), c
iting
Rea
dio
(192
7),
a fa
lse re
cord
per
haps
bas
ed o
n m
isint
erpr
etat
on o
f con
fusin
g te
xt
in R
yckm
an &
Ben
tley
(197
9, p
. 36)
.
Hor
tens
ia si
mili
s (W
alke
r, 18
51)
Cic
adel
linae
Trin
idad
Mye
rs (1
929,
p.4
74) (
as K
olla
sim
ilis)
repo
rtin
g co
mm
unic
atio
n w
ith H
.A. B
allo
u.
Hor
tens
ia si
mili
s (W
alke
r, 18
51)
Cic
adel
linae
Ryck
man
& B
entle
y (1
979,
p.3
3) (a
s Kol
la si
mili
s), c
iting
Mye
rs
(192
9).
(Con
tinue
d )
Tabl
e 1.
(C
ont.)
-
214 E.Domínguez Núñez and A. Aiello / Terrestrial Arthropod Reviews 6 (2013) 201–225
Typh
locy
ba q
uerc
us (F
abric
ius,
1777
), bu
t M
yers
had
dou
bts a
bout
the
iden
tifica
tion
Typh
locy
bina
eJa
pan
Mye
rs (1
929,
p.4
75) [
He
had
doub
ts ab
out t
he id
entifi
catio
n],
repo
rtin
g un
publ
ished
obs
erva
tion
by T
. Esa
ki.
Typh
locy
ba q
uerc
us (F
abric
ius,
1777
)Ty
phlo
cybi
nae
Ryck
man
& B
entle
y (1
979,
p.3
3), c
iting
Mye
rs (1
929)
, but
Mye
rs
had
doub
ts ab
out t
he id
entifi
catio
n.
Erat
oneu
ra m
acul
ata
(Gill
ette
, 189
8)Ty
phlo
cybi
nae
Japa
nM
yers
(192
9, p
.475
) [as
Ery
thro
neur
a (Z
ygin
a) m
acul
ata?
], re
port
ing
unpu
blish
ed o
bser
vatio
n by
T. E
saki
.
Erat
oneu
ra m
acul
ata
(Gill
ette
, 189
8)Ty
phlo
cybi
nae
Ryck
man
& B
entle
y (1
979,
p.3
3) (a
s Ery
thro
neur
a m
acul
ata)
, ci
ting
Mye
rs (1
929)
.
Empo
asca
?Ty
phlo
cybi
nae
no lo
calit
y gi
ven,
bu
t lik
ely
to h
ave
been
Cin
nam
ara,
As
sam
, Ind
ia
Andr
ews (
1929
, p.9
6), p
erso
nal e
xper
ienc
e re
port
ed in
lette
r rea
d by
E.B
. Pou
lton
at 1
6 O
ct. 1
929
mee
ting
of th
e En
tom
olog
ical
So
ciet
y of
Lon
don.
Kyb
os sm
arag
dula
(Dla
bola
, 195
8c:5
1)Ty
phlo
cybi
nae
Engl
and
Mye
rs (1
929,
p.4
75 &
477
), re
port
ing
the
as y
et u
npub
lishe
d ex
perie
nce
of W
.E. C
hina
.
Kyb
os sm
arag
dula
(Dla
bola
, 195
8c:5
1)Ty
phlo
cybi
nae
Esth
er, S
urre
y [E
ngla
nd]
Chi
na (1
929,
p.9
6-97
) (as
Em
poas
ca sm
arag
dula
), hi
s per
sona
l ex
perie
nce
repo
rted
in n
ote
read
by
E.B.
Pou
lton
at 1
6 O
ct. 1
929
mee
ting
of th
e En
tom
olog
ical
Soc
iety
of L
ondo
n, w
here
in h
e m
entio
ns th
at “
Mye
rs h
as re
cent
ly p
ublis
hed
a no
te o
n th
is su
bjec
t, bu
t I h
ave
had
no ti
me
to tr
ace
it.”
Kyb
os sm
arag
dula
(Dla
bola
, 195
8c:5
1)Ty
phlo
cybi
nae
Ryck
man
& B
entle
y (1
979,
p.3
3), c
iting
Mye
rs (1
929)
.
Neo
alitu
rus t
enell
us (E
mel
jano
v, 19
99c:
56)
Del
toce
phal
inae
Uni
vers
ity o
f C
alifo
rnia
[USA
]U
singe
r (19
34, p
.97)
(as E
utet
tix te
nellu
s), re
port
ing
unpu
blish
ed
expe
rienc
e of
Juliu
s Fre
itag.
Tabl
e 1.
(C
ont.)
(Con
tinue
d )
-
E.Domínguez Núñez and A. Aiello / Terrestrial Arthropod Reviews 6 (2013) 201–225 215
Neo
alitu
rus t
enell
us (E
mel
jano
v, 19
99c:
56)
Del
toce
phal
inae
Her
ms (
1950
, p.5
74) (
as E
utet
tix te
nellu
s), c
iting
Usin
ger (
1934
).
Neo
alitu
rus t
enell
us (E
mel
jano
v, 19
99c:
56)
Del
toce
phal
inae
Ryck
man
& B
entle
y (1
979,
p.4
2) (a
s Eut
ettix
tene
llus),
citi
ng
Usin
ger (
1934
).N
eoal
ituru
s ten
ellus
(Em
elja
nov,
1999
c:56
)D
elto
ceph
alin
aeC
alifo
rnia
[USA
]Al
exan
der (
1984
, p.5
8) (a
s Eut
ettix
tene
llus),
citi
ng U
singe
r (19
34).
Eryt
hron
eura
com
es (S
ay, 1
825)
Typh
locy
bina
eFr
esno
, Cal
iforn
ia
[USA
]U
singe
r (19
34, p
.97-
98),
repo
rtin
g un
publ
ished
exp
erie
nces
of
Cal
vert
E. N
orla
nd a
nd G
ordo
n L.
Sm
ith.
Eryt
hron
eura
com
es (S
ay, 1
825)
Typh
locy
bina
eH
erm
s (19
50, p
.574
), ci
ting
Usin
ger (
1934
).Er
ythr
oneu
ra co
mes
(Say
, 182
5)Ty
phlo
cybi
nae
Ryck
man
& B
entle
y (1
979,
p.4
2) (a
s Ery
thro
neur
a co
mes
), ci
ting
Usin
ger (
1934
).Er
ythr
oneu
ra co
mes
(Say
, 182
5)Ty
phlo
cybi
nae
Cal
iforn
ia [U
SA]
Alex
ande
r (19
84, p
.58)
(miss
pelle
d as
Ery
thro
nem
a co
mes)
, citi
ng
Usin
ger (
1984
).
Exiti
anus
fasci
olat
us (M
elic
har,
1911
)D
elto
ceph
alin
aeIn
Sal
ah [s
outh
ern
Alge
ria]
Fole
y &
Aud
ouze
(193
8, p
.526
) (as
Ath
ysan
us v
ulne
rans
), re
port
ing
obse
rvat
ions
by
Fonq
ue in
Sep
t. 19
24.
Exiti
anus
fasci
olat
us (M
elic
har,
1911
)D
elto
ceph
alin
aesu
d al
gerie
nBr
umpt
(194
9, p
.127
6) (a
s Ath
ysan
us v
ulne
rans
), re
port
ing
obse
rvat
ion
mad
e by
Dr.
Fouq
ue (m
isspe
lled
as F
onqu
e).
Neo
alitu
rus t
enell
us (E
mel
jano
v, 19
99c:
56)
Del
toce
phal
inae
Loga
n, U
tah
[USA
]K
now
lton
(195
1, p
.112
) (as
Circ
ulife
r ten
ellus
), ob
serv
atio
ns b
y hi
m a
nd M
.F. B
owen
.
Eryt
hron
eura
Typh
locy
bina
eW
ad M
edan
i [Su
dan]
Lew
is (1
958,
p.4
5), a
ppar
ently
an
inde
pend
ent r
epor
t.
Eryt
hron
eura
Typh
locy
bina
eSu
dan
Ryck
man
& B
entle
y (1
979,
p.2
8), c
iting
Lew
is (1
958)
.
Delt
ocep
halu
sD
elto
ceph
alin
aeW
ad M
edan
i [S
udan
]Le
wis
(195
8, p
.45)
, app
aren
tly a
n in
depe
nden
t rep
ort.
Delt
ocep
halu
sD
elto
ceph
alin
aeSu
dan
Ryck
man
& B
entle
y (1
979,
p.2
8), c
iting
Lew
is (1
958)
.
(Con
tinue
d )
Tabl
e 1.
(C
ont.)
-
216 E.Domínguez Núñez and A. Aiello / Terrestrial Arthropod Reviews 6 (2013) 201–225
Exiti
anus
fasci
olat
us (M
elic
har,
1911
)D
elto
ceph
alin
aeK
hart
oum
[Sud
an]
Lew
is (1
958,
p.4
5) (a
s E. v
ulne
rans
Ber
gevi
n). Th
ere
is co
nfus
ion
here
: the
gen
us n
ame
is no
t spe
lled
out a
nd th
e pr
eced
ing
genu
s is
Eusce
lis, a
lthou
gh v
ulne
rans
Ber
gevi
n w
as d
escr
ibed
in A
thys
anus
. Re
gard
less
, app
aren
tly th
is is
an in
depe
nden
t rep
ort b
ecau
se a
ll pr
evio
us re
cord
s are
for S
alah
.
Exiti
anus
fasci
olat
us (M
elic
har,
1911
)D
elto
ceph
alin
aeSu
dan
Ryck
man
& B
entle
y (1
979,
p.2
8) (a
s E. v
ulne
rans
), ci
ting
Lew
is (1
958)
.
Exiti
anus
?D
elto
ceph
alin
aeW
ad M
edan
i [S
udan
]Le
wis
(195
8, p
.45)
, app
aren
tly a
n in
depe
nden
t rep
ort.
Tham
note
ttix
2 sp
p.D
elto
ceph
alin
aeW
ad M
edan
i [S
udan
]Le
wis
(195
8, p
.46)
, app
aren
tly a
n in
depe
nden
t rep
ort.
Empo
asca
dec
ipien
s (Pa
oli,
1930
) or
Empo
asca
sola
ni (C
urtis
, 184
6)Ty
phlo
cybi
nae
Oxf
ord
[Eng
land
]M
cCra
e (1
974,
238
). Sp
ecim
en d
eter
min
ed b
y W
. J. L
e Q
uesn
e as
as
a fe
mal
e of
eith
er E
mpo
asca
dec
ipien
s or E
. pte
ridis,
the
fem
ales
of
whi
ch “c
anno
t be
diffe
rent
iate
d.”
Hor
tens
ia si
mili
s (W
alke
r, 18
51)
Cic
adel
linae
Cub
aN
ovoa
& A
layo
(198
6, p
.80)
.
Hor
tens
ia si
mili
s (W
alke
r, 18
51)
Cic
adel
linae
Repu
blic
of P
anam
aG
uerr
a V.
(200
2, p
.38-
39),
unpu
blish
ed th
esis.
Hom
alod
isca
vitri
penn
is G
erm
ar, 1
821
Cic
adel
linae
Fren
ch P
olyn
esia
Gra
ndgi
rard
et a
l. (2
006,
p.4
35) (
as H
omal
odisc
a co
agul
ata)
.
Tabl
e 1.
(C
ont.)
-
E.Domínguez Núñez and A. Aiello / Terrestrial Arthropod Reviews 6 (2013) 201–225 217
we surmise that only 37 are original, and the rest are citation upon citation of those 37. Not included in the 174 are three false records that were based upon a misreading of Readio (1927).
We add 19 new records, all from Panama, representing 13 genera, and at least 13 species to the list of Cicadellidae known to have attacked humans (Table 2). These new records represent five subfamilies, two of which, Gyponinae and Xestocephalinae, have not been reported previously. Eleven are first records for their genera: Dilobopterus, Tylozygus, Balclutha, Chlorotettix, Hecalapona, Planicephalus, Plesiommata, Portanus, Xestocephalus, Xyphon, and Zyginama. Two attacks were to the back of the neck, but the other 17 were to the extremities, mostly the forearm, and in each case caused itching or mild stinging lasting several seconds to several minutes.
Fifteen of the attacks took place at night or at dusk, usually on people reading or watching television. One of the four daytime attacks was by an individual of Dilobopterus stolli that alighted on EDN’s leg and probed repeatedly for some three minutes but did not pierce the skin. It was then captured in a vial and placed on the leg of EDN’s wife (LV). The insect immediately pierced her skin, causing pain and itching. A second incident involved a lab researcher who was attacked upon reaching into a brightly lighted chamber containing his study subjects, Hortensia similis (Guerra, 2002). A third daytime attacker was a typhlocybine that alighted on EDN while he was wait-ing at a bus stop. The fourth daytime attack took place in a patio. A fifth daytime attack is not included in our data because the specific insect individual was not cap-tured; it was to a person (LV) who was watering a garden, and thus disturbed a large number of Dilobopterus stolli on their plant, Pseuderanthemum reticulatum (Hooker f.) Radlkofer (Acanthaceae). Including our own contributions, we calculate that now there are 193 published records of Cicadellidae having bitten humans, of which 56 are the original observations, and that they represent 30 genera and at least 39 species in five subfamilies.
In addition to our experiences, Pierre Jolivet has been probed by various species near rice fields and the Awash River in Ethiopia; and by Nephotettix, probably virescens, dur-ing a typhoon in Hong-Kong (personal communication). As well, William Eberhard (in the Neotropics) has been probed occasionally by leafhoppers (personal communi-cation). Neither of them captured their attackers.
Discussion
Several authors doubted that leafhoppers showed interest in humans until they experi-enced it themselves. Donovan (1920a, 1920b) commented “Several people used to complain to me of being stung by these insects, but, not being cognizant at the time of such habits among the Homoptera, I discountenanced these tales of the presumed aggressors. It was only when I was myself a victim of the insects’ action that I was disillusioned.” Lawson (1920) stated “The writer has had several people tell him about being bitten by little green leafhoppers, but not till a short time ago did he have any personal proof of the fact. One night, while collecting under a light, he felt a little prick on his hand, and on looking down saw a little green leafhopper [Empoasca fabae
-
218 E.Domínguez Núñez and A. Aiello / Terrestrial Arthropod Reviews 6 (2013) 201–225Ta
ble
2. C
icad
ellid
ae th
at h
ave
prob
ed h
uman
skin
in th
e Re
publ
ic o
f Pan
ama
Nam
eSu
bfam
ilyG
ende
rD
ay/n
ight
Dat
eLo
calit
y, vi
ctim
, and
par
t of b
ody
atta
cked
Dep
ositi
on
Dilo
bopt
erus
stol
li Si
gnor
et, 1
850
Cic
adel
linae
mal
eD
2011
Nov
. 13
Pana
má:
San
Isid
ro, E
l Val
le, i
n pa
tio: p
robe
d ED
N’s
leg
with
no
succ
ess,
but p
robe
d Li
nette
Veg
a’s (L
V) l
eg
afte
r bei
ng tr
ansfe
rred
ther
e
MIU
P
Hor
tens
ia si
mili
s (W
alke
r, 18
51)
Cic
adel
linae
fem
ale
N20
09 D
ec. 2
3Pa
nam
á: S
an Is
idro
, El V
alle
, in
livin
g ro
om: p
robe
d LV
’s fo
rear
mM
IUP
Hor
tens
ia si
mili
s (W
alke
r, 18
51)
Cic
adel
linae
fem
ale
D20
01Pa
nam
á: U
nive
rsid
ad d
e Pa
nam
á, in
Lab
. de
Estu
dios
Bi
ológ
icos
de
Plag
as A
gríc
olas
, in
lab:
“atta
cked
” Is
mae
l Gue
rra
[Unp
ublis
hed
thes
is]
MIU
P
Tylo
zygu
s geo
met
ricus
(S
igno
ret,
1854
)C
icad
ellin
aefe
mal
eN
2008
Oct
. 23
Pana
má:
Arr
aijá
n, L
oma
del R
ío, 8
.940
7 -7
9.65
68, i
n liv
ing
room
: pro
bed
AA’s
fore
arm
STR
I
Xyph
on re
ticul
atum
(S
igno
ret 1
854)
Cic
adel
linae
mal
edu
sk20
12 F
eb. 4
Pana
má:
Can
al A
rea,
nea
r Can
al A
dmin
istra
tion
Build
ing:
pro
bed
LV’s
fore
arm
MIU
P
Plesi
omm
ata
sp.
Cic
adel
linae
mal
edu
sk20
12 N
ov. 1
5Pa
nam
á: C
hiriq
uí, D
avid
, Dol
egui
ta, 8
th S
treet
: pr
obed
Indi
ra S
imon
’s le
gST
RI
Plesi
omm
ata
sp.
Cic
adel
linae
fem
ale
N20
12 N
ov. 2
9Pa
nam
á: A
rrai
ján,
Pue
rta
de H
ierr
o: p
robe
d Je
rald
ín
Verg
ara’s
left
ring
finge
rM
IUP
Balcl
utha
sp.
Del
toce
phal
inae
fem
ale
N20
09 O
ct. 2
6Pa
nam
á: S
an Is
idro
, El V
alle
, in
livin
g ro
om: p
robe
d LV
’s fo
rear
mM
IUP
Chl
orot
ettix
min
imus
Ba
ker,
1898
Del
toce
phal
inae
mal
eN
2012
Mar
. 14
Pana
má:
Arr
aijá
n, L
oma
del R
ío, 8
.940
7 -7
9.65
68,
unde
r rea
ding
lam
p: p
robe
d th
e ba
ck o
f AA’
s han
d (0
2: 5
2 H
)
STR
I
Plan
iceph
alus
flav
icosta
(S
tål,
1862
)D
elto
ceph
alin
aem
ale
N20
12 M
ar. 5
Pana
má:
San
Isid
ro, E
l Val
le, i
n liv
ing
room
: pro
bed
LV’s
fore
arm
MIU
P
Hec
alap
ona
brev
isens
D
eLon
g an
d Fr
eyta
g,
1975
Gyp
onin
aefe
mal
eN
2008
Aug
. 19
Pana
má:
Arr
aijá
n, L
oma
del R
ío, 8
.940
7 -7
9.65
68,
unde
r rea
ding
lam
p: p
robe
d AA
’s fo
rea
![The New Zealand Aleyrodidae (Hemiptera: Homoptera)€¦ · The New Zealand Aleyrodidae (Hemiptera: Homoptera)1 1. ]. DUMBLETON2 THE PIONEER WORKER on the New Zealand Aleyrodidae was](https://static.fdocuments.us/doc/165x107/5eb97aadf345c43a4770ccd8/the-new-zealand-aleyrodidae-hemiptera-homoptera-the-new-zealand-aleyrodidae-hemiptera.jpg)
